Host records and geographical distribution of Corynosoma magdaleni, C. semerme and C. strumosum (Acanthocephala: Polymorphidae).

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Host records and geographical distribution of

Corynosoma magdaleni, C. semerme and C.

strumosum (Acanthocephala: Polymorphidae)

Sonja Leidenberger

, Sven Boström

, Matthew Thomas Wayland

‡ School of Bioscience, Department of Biology and Bioinformatics, University of Skövde, Skövde, Sweden § Swedish Museum of Natural History, Department of Zoology, Stockholm, Sweden

| University of Cambridge, Cambridge, United Kingdom

Corresponding author: Sonja Leidenberger (sonja.leidenberger@his.se) Academic editor: Yasen Mutafchiev

Received: 27 Jan 2020 | Accepted: 30 Mar 2020 | Published: 06 Apr 2020

Citation: Leidenberger S, Boström S, Wayland MT (2020) Host records and geographical distribution of Corynosoma magdaleni, C. semerme and C. strumosum (Acanthocephala: Polymorphidae). Biodiversity Data Journal 8: e50500. https://doi.org/10.3897/BDJ.8.e50500

Abstract

A literature survey was conducted to investigate the host and geographical distribution

patterns of three

Corynosoma

species (Acanthocephala: Polymorphidae),

viz. C.

magdaleni

,

C. semerme

and

C. strumosum

. All three species appear to be restricted to the

Northern Hemisphere. Occurrence records of

C. magdaleni

are limited to the Northern

Atlantic coasts, while

C. semerme

has a circumpolar distribution. The geographical range

of

Corynosoma strumosum

encompasses the distributions of the other two species, but

also extends into warmer southern regions. Some

Corynosoma

populations are living with

their definitive hosts in very isolated locations, such as in the brackish Baltic Sea or

different freshwater lakes (e.g. Lake Saimaa). All three species have a heteroxenous life

cycle, comprising a peracaridan intermediate host, a fish paratenic host and a mammalian

definitive host. Occasionally, an acanthocephalan may enter an accidental host, from which

it is unable to complete its life cycle. The host records reported here are categorised by

type,

i.e.

intermediate, paratenic, definitive or accidental. While most of the definitive hosts

are shared amongst the three

Corynosoma

species,

C. strumosum

showed the broadest

range of paratenic hosts, which reflects its more extensive geographical distribution. One

aim of this study and extensive literature summary is to guide future sampling efforts and

‡ § |

© Leidenberger S et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

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therewith contribute to throw more light on the on-going species and morphotype

discussion for this interesting parasite species.

Keywords

Acanthocephala,

Corynosoma

, host record, life cycle, Northern Hemisphere, seal and

zoogeography

Introduction

The genus

Corynosoma

comprises 43 marine species (Amin 2013), that infect mammals

and piscivorous birds. Aznar et al. (2006) showed that the original concept of

Corynosoma

was not a monophyletic genus. Phylogenetic analysis revealed that the marine

Corynosoma

species were more closely related to the genus

Andracantha

Schmidt, 1975

than to the species living in freshwater habitats. Aznar et al. (2006) erected the genus

Pseudocorynosoma

Aznar, Pérez-Ponce de León and Raga, 2006 for the freshwater

species. Recent phylogenetic analysis based on ribosomal ITS1-5.8S-ITS2 and COI

sequences showed that

Corynosoma

spp., hosted by Pinnipedia and marine Mustelidae

formed a monophyletic group (Waindok et al. 2018).

The systematics of

Corynosoma

species from seals in northern Europe have recently been

investigated using both molecular markers (Waindok et al. 2018) and morphology

(Leidenberger et al. 2019). These studies have highlighted the limitations of using

traditional morphological characters alone for diagnosing species. To determine the true

species diversity in this genus, tandem morphological and molecular studies will be

required. To guide future sampling efforts, we have conducted a comprehensive literature

survey of the host and geographical distribution of three commonly encountered taxa:

C.

magdaleni

Montreuil, 1958,

C. semerme

(Forssell, 1904) Lühe, 1911 and

C. strumosum

(Rudolphi, 1802) Lühe, 1904.

Material and methods

Host and geographical records

Host and geographical records for the three acanthocephalan taxa were collected from

literature. Manual searches were conducted using Pubmed, Web of Science and Google

Scholar. Additionally, we used the R (R Core Team 2019) package helminthR (Dallas 2016)

to extract records from the Host-Parasite Database of The Natural History Museum in

London (Gibson et al. 2005). Geographical records for each

Corynosoma

species were

summarised by MEOW ecoregions (Spalding et al. 2007) and then plotted as a distribution

map using the meow R-package (Byrnes 2016).

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For species names, we followed the nomenclature given by the Catalogue of Life (2019

Annual Checklist: Roskov et al. 2019) and WoRMS (WoRMS Editorial Board 2019).

The term

accidental host

was used here for a host, where the parasite usually is not found,

because the host is not suitable for the parasite's development. This can mean that the

accidental host also becomes a dead-end, because the life cycle of the parasite is blocked

(e.g. no observed gravid female and/or mature individuals) and the parasite does not reach

its definitive host. Since in many literature references, this detailed information was not

given, we used the term accidental host as generic term for all of the dead-end/accidental/

incidental host records we found.

Results and discussion

Zoogeography

Our literature survey on the zoogeography showed that the three species of

Corynosoma

(

C.

magdaleni

,

C. semerme

and

C. strumosum

) and their definitive hosts are restricted to

the Northern Hemisphere (Figs 1, 2, 3). The relatively few records for

C. magdaleni

suggest that this taxon is restricted to the North Sea, Baltic Sea and northwest Atlantic. By

contrast,

C. semerme

has a circumpolar distribution, having been reported from the Arctic

Ocean, north Atlantic and north Pacific (e.g. Popov and Fortunato 1987, Kaimoto et al.

2018, Waindok et al. 2018) (Fig. 2). The geographical range of

C. strumosum

encompasses the distributions of the other two taxa, but also extends into more southern

regions, such as the Mediterranean Sea and coast of California (Fig. 3).

Figure 1.

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In most of the Baltic Sea studies,

C. semerme

was the species most commonly found in

seals, followed by

C. magdaleni

and only rare infections were found for

C. strumosum

(Nickol et al. 2002, Sinisalo and Valtonen 2003, Leidenberger and Bäcklin 2008, Waindok

et al. 2018). Co-infections have been observed (Nickol et al. 2002, Leidenberger and

Bäcklin 2008, Leidenberger et al. 2019), most commonly involving

C. semerme

and

C.

magdaleni

(Sinisalo and Valtonen 2003, Leidenberger and Bäcklin 2008, Waindok et al.

2018). Interestingly, land-locked seal species show no mixed infections, but only single

infections, for example,

Phoca hispida saimensis

in Lake Saimaa only by

C. magdaleni

Figure 2.

Geographical distribution of

Corynosoma semerme

.

Figure 3.

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found in the Caspian Sea is

C. caspicum

and not

C. strumosum

. Indeed,

C. strumosum

from the land-locked Caspian seal recorded by Amin et al. 2011 was characterised with

proboscis hooks and trunk spines distinct from other Northern European morphological

descriptions (Nickol et al. 2002, Waindok et al. 2018, Leidenberger et al. 2019), suggesting

that they may not be conspecific. Surprisingly,

Corynosoma caspicum

Golvan and

Mokhayer, 1973, described based on immature specimens from sturgeons in Caspian Sea

(Golvan and Mokhayer 1973) and often found in three-spined stickleback (

Gasterosteus

aculeatus

) (Niksirat et al. 2006), has not yet been reported in

Pusa caspica

or another

definitive host. Waindok et al. (2018) described some individuals of

C. strumosum

from the

German North and Baltic Seas with different proboscis morphology, similar to those

described by Amin et al. (2011) and stated this as

C. magaleni

isolate Pv1NS instead.

Additionally, they found a cryptic species refered to as "Candidatus

Corynosoma nortmeri

sp. nov."' Waindok et al. (2018) in the European study area.

The extent of the geographical and host ranges of the three taxa correlates with the year of

their description.

Corynosoma strumosum

was the first species of the genus to be

described (Rudolphi 1802) and has, by far, the broadest geographical and host

distributions.

Corynosoma semerme

, described by Forssell (1904), appears to have a

slightly more limited geographical range and far fewer paratenic host records. Relatively

few hosts have been reported for

C. magdaleni

, described by Montreuil (1958) and its

geographical range appears to be restricted to the northern Atlantic, in contrast to the

circumpolar distributions of the other two species. Up until its description in 1958,

specimens of

C. magdaleni

were almost certainly assigned to the morphologically similar

C. strumosum.

The potential bias resulting from taxon age should be kept in mind when

comparing the reported host and geographical distributions of the three taxa (Fig. 3).

Overview of the life cycle

The heteroxenous life cycle of

Corynosoma

species involves a peracaridan intermediate

host, a paratenic host (fish) and a mammalian definitive host. While there are numerous

studies on

C. semerme

and

C. strumosum

(Tables 3, 4, 5, 6, 7, 8), there have been few

studies on

C. magdaleni

, besides its original description by Montreuil 1958 (Tables 1, 2).

The intermediate host of

C. magdaleni

is unknown. Lake Saimaa may be a good location

to search for the intermediate host of

C. magdaleni

, because the other

Corynosoma

species appear to be absent from this land-locked waterbody (Sinisalo and Valtonen 2003,

Sinisalo et al. 2003). A total of five species of paratenic host (all Actinopterygii) have been

reported for

C. magdaleni

; three from Canada and two from the Baltic Sea (Table 1). In

Canada, only

C. magdaleni

is reported together with

C. wegeneri

Heinze, 1934. In the

definitive hosts (Table 2), the species seems to prefer the last part of the small intestine

(ileum) and the colon.

Corynosoma semerme

has been reported from the whole intestine,

but is typically found at a higher density in the large intestine, especially the cecum and

rectum (Nickol et al. 2002, Leidenberger and Bäcklin 2008), where

C. strumosum

and

C.

magdaleni

are not generally observed. Microhabitat segregation might facilitate

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reproductive isolation of the three species, especially in mixed infections that are often

observed in seals from the Baltic Sea (Nickol et al. 2002, Sinisalo and Valtonen 2003,

Leidenberger and Bäcklin 2008), but less commonly in the North Sea (Waindok et al.

2018). While the three

Corynosoma

species share most of their definitive hosts, they show

some differences in their paratenic hosts. We found that

C. strumosum

shares three

paratenic hosts with both other species, none with

C. magdaleni

only, but > 20 paratenic

hosts are shared with

C. semerme.

More than 80 paratenic hosts were described for

C.

strumosum

and these are, up to now, not reported from any of the other two species

(Tables 1, 4, 6).

Host order Host family Host species Host

vernacular name

Geographical locality

Reference(s)

Perciformes Percidae Gymnocephalus cernua (Linnaeus, 1758) (as Gymnocephalus cernuus)

ruffe Baltic Sea,

North-eastern Bothnian Bay

Valtonen et al. 2001

Pleuronectiformes Pleuronectidae Hippoglossus hippoglossus (Linnaeus, 1758)

Atlantic halibut Atlantic, Canada Arai 1989

Pleuronectiformes Pleuronectidae Hippoglossus hippoglossus (Linnaeus, 1758)

Atlantic halibut Magdalen Islands region of the Gulf of St. Lawrence

Montreuil 1958

Salmoniformes Salmonidae Salvelinus fontinalis (Mitchill, 1814)

brook charr Tabusintac River, New Brunswick

Frimeth 1987

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758)

fourhorn sculpin Baltic Sea, Bothnian Bay

Sinisalo and Valtonen 2003

Scorpaeniformes Cottidae Myoxocephalus scorpius (Linnaeus, 1758)

shorthorn sculpin Atlantic, Canada Arai 1989

Scorpaeniformes Cottidae Myoxocephalus scorpius (Linnaeus, 1758)

shorthorn sculpin Magdalen Islands region of the Gulf of St. Lawrence Montreuil 1958

Table 1.

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Host order

Host family

Host species Host

vernacular name

Geographical locality Reference

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Baltic Sea Waindok et al. 2018

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Baltic Sea, Baltic Proper, Bothnian Sea, Bothnian Bay, Sweden

Leidenberger and Bäcklin 2008

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Baltic Sea, Finland, Gulf of Bothnia, Åland Island

Nickol et al. 2002

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Baltic Sea, Sweden Leidenberger et al. 2019

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Eastern Canada Montreuil 1958

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Eastern Canada Montreuil 1958

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal North Sea, Schleswig-Holstein

Waindok et al. 2018

Carnivora Phocidae Phoca vitulina Linnaeus, 1758 (as Phoca vitulina concolor De Kay, 1842)

harbour seal Eastern Canada Montreuil 1958

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Baltic Sea Leidenberger et al. 2019

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Baltic Sea, Germany, Schleswig-Holstein

Waindok et al. 2018

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Baltic Sea, Bothnian Bay

Valtonen et al. 2004

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Baltic Sea, Gulf of Finland

Delyamure et al. 1980

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal European Arctic Treshchev 1982* citet by Raga 1992

Carnivora.

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Host order

Host family

Host species Host

vernacular name

Geographical locality Reference

Carnivora Phocidae Pusa hispida saimensis Nordquist, 1899

Saimaa ringed seal

Lake Saimaa, Finland Sinisalo et al. 2004, Aznar et al. 2006, García-Varela et al. 2011

Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Artiodactyla Monodontidae Delphinapterus leucas (Pallas, 1776)

Beluga, Beluga whale, white whale

Bristol Bay, Alaska, Bering Sea

Neiland 1962, Measures et al. 1995 Carnivora Odobenidae Odobenus rosmarus

(Linnaeus, 1758)

walrus Arctic Ocean,

Northern Atlantic

Delyamure 1961, von Sprehn 1966 cited by Raga 1992 Carnivora Otariidae Callorhinus ursinus

(Linnaeus, 1758)

Northern fur seal Alaska, Bering Sea, St. Paul Island

Kuzmina et al. 2012

Carnivora Otariidae Callorhinus ursinus (Linnaeus, 1758) (as Callorhinus alascanus)

Northern fur seal North America Van Cleave 1953

Carnivora Phocidae Cystophora cristata (Erxleben, 1777)

hooded seal European Arctic Delyamure 1961*, Treshchev 1982* cited by Raga 1992 Carnivora Phocidae Erignathus barbatus

(Erxleben, 1777)

bearded seal North America Van Cleave 1953

Carnivora Phocidae Erignathus barbatus

(Erxleben, 1777) bearded seal White Sea Popova et al. 1975* cited by Raga 1992 Carnivora Phocidae Halichoerus grypus

(Fabricius, 1791)

grey seal Arctic Ocean, Northern Atlantic, European Arctic

von Sprehn 1966* cited by Raga 1992

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Germany Lühe 1911

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Germany Aznar et al. 2006

Table 3.

Definitive hosts of

Corynosoma semerme

. *Source not viewed, ^Found in Host-Parasite Database

NHM London without further reference details.

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Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Baltic Sea, Finland, Gulf of Bothnia, Åland Island

Nickol et al. 2002

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Baltic Sea, Baltic Proper, Bothnian Sea, Bothnian Bay, Sweden

Leidenberger and Bäcklin 2008

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Baltic Sea, Germany, Schleswig-Holstein

Waindok et al. 2018

Carnivora Phocidae Histriophoca fasciata (Zimmerman, 1783) (as Phoca fasciata)

ribbon seal Bering Sea, Alaska Rausch et al. 1990

Carnivora Phocidae Histriophoca fasciata (Zimmerman, 1783) (as Phoca fasciata)

ribbon seal Arctic Ocean, Bering Sea

Shults and Frost 1988

Carnivora Phocidae Histriophoca fasciata (Zimmerman, 1783) (as Phoca fasciata)

ribbon seal Sea of Okhotsk Shults and Frost 1988

Carnivora Phocidae Pagophilus groenlandicus (Erxleben, 1777) (as Phoca groenlandica) harp seal, Greenland seal Arctic Ocean, Northern Atlantic, European Arctic von Sprehn 1966* cited by Raga 1992

Carnivora Phocidae Phoca largha Pallas, 1811

spotted seal, larga seal

Sea of Okhotsk Sasaki et al. 2019

Carnivora Phocidae Phoca largha Pallas, 1811

spotted seal, larga seal

Russia Aznar et al. 2006

Carnivora Phocidae Phoca largha Pallas, 1811

spotted seal, larga seal

Bering Sea, Alaska Rausch et al. 1990

Carnivora Phocidae Phoca largha Pallas, 1811

spotted seal, larga seal

Arctic Ocean, Bering Sea

[On the helminthofauna of largi and harbour seals.]^ Carnivora Phocidae Phoca largha Pallas,

1811

spotted seal, larga seal

Sea of Okhotsk [On the helminthofauna of largi and harbour seals.]^

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Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Germany Lühe 1911

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Sweden Lundström 1942

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal North Sea, Germany, Schleswig-Holstein

Waindok et al. 2018

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Arctic Ocean, Northern Atlantic, European Arctic

von Sprehn 1966* cited by Raga 1992

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Alaska, Bering Sea, St. Paul Island

Kuzmina et al. 2012

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Germany Lühe 1911

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Baltic Sea, Bothnian Bay, Finland

Helle and Valtonen 1981

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Baltic Sea, Bothnian Bay, Finland

Valtonen and Helle 1988

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Baltic Sea, Bothnian Bay, Finland

Valtonen et al. 2004

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal North America Van Cleave 1953

Carnivora Phocidae Pusa hispida (Schreber, 1775) (as Phoca foetida)

ringed seal Baltic Sea, Sweden, Östra Rönnskär

Forssell 1904

Carnivora Phocidae Pusa hispida (Schreber, 1775) (as Phoca foetida)

ringed seal Baltic Sea, Finland, Tvärminne

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Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Acipenseriformes Acipenseridae Acipenser sturio Linnaeus, 1758

sturgeon USSR (CIS) Shulman

1954* Acipenseriformes Acipenseridae Acipenser sturio

Linnaeus, 1758

sturgeon USSR (CIS) Shulman

1954* Anguilliformes Anguillidae Anguilla anguilla

(Linnaeus, 1758)

European eel Sweden Lundström

1942 Anguilliformes Anguillidae Anguilla anguilla

(Linnaeus, 1758)

European eel Europe Van Cleave 1953 Clupeiformes Clupeidae Clupea harengus

Linnaeus, 1758

herring Sweden Lundström

1942 Clupeiformes Clupeidae Clupea harengus

Linnaeus, 1758 (as Clupea harengus membras)

herring Baltic Sea, Finland, Tvärminne

Forssell 1904

Clupeiformes Clupeidae Clupea harengus Linnaeus, 1758 (as Clupea harengus membras)

herring Baltic Sea, Finland, Tvärminne

Forssell 1905

Clupeiformes Clupeidae Clupea harengus Linnaeus, 1758 (as Clupea harengus membras)

herring Germany Lühe 1911

Clupeiformes Clupeidae Clupea harengus Linnaeus, 1758 (as Clupea harengus membras)

herring Europe Van Cleave

1953

Cypriniformes Cyprinidae Blicca bjoerkna (Linnaeus, 1758)

white bream Sweden Lundström

1942 Cypriniformes Cyprinidae Blicca bjoerkna

(Linnaeus, 1758)

white bream Europe Van Cleave

1953 Cypriniformes Cyprinidae Pelecus cultratus

(Linnaeus, 1758) ziege Finno-Karelian ASSR Rumyantsev and Ieshko 1997*

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Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Esociformes Esocidae Esox lucius Linnaeus, 1758

pike Baltic Sea,

Finland, Tvärminne

Forssell 1905

Gadiformes Gadidae Boreogadus saida (Lepechin, 1774)

polar cod Arctic Ocean, USSR zone, Pechora Sea

Karasev and Mitenev 1993* Gadiformes Gadidae Gadus morhua

Linnaeus, 1758

cod Baltic Sea,

Finland, Tvärminne

Forssell 1905

Gadiformes Gadidae Gadus morhua Linnaeus, 1758 (as Gadus callarias)

cod Sweden Lundström

1942

Gadiformes Gadidae Gadus morhua Linnaeus, 1758 (as Gadus callarias)

cod Europe Van Cleave

1953

Gadiformes Lotidae Enchelyopus cimbrius (Linnaeus, 1766) (as Onos cimbrius)

fourbeard rockling Europe Van Cleave 1953

Gadiformes Lotidae Lota lota (Linnaeus, 1758)

burbot Baltic Sea,

Bothnian Bay

Valtonen and Julkunen 1995 Gadiformes Lotidae Lota lota (Linnaeus,

1758) (as Lota vulgaris)

burbot Sweden Lundström

1942 Gadiformes Lotidae Lota lota (Linnaeus,

1758) (as Lota vulgaris)

burbot Europe Van Cleave

1953 Gadiformes Lotidae Lota lota (Linnaeus,

1758) burbot Tuluksak, Bering Sea, Alaska Rausch et al. 1990

Gasterosteiformes Gasterosteidae Gasterosteus aculeatus Linnaeus, 1758

three-spined stickleback

Sweden Lundström

1942 Lophiiformes Lophiidae Lophius piscatorius

Linnaeus, 1758

angler Sweden Lundström

1942 Osmeriformes Osmeridae Osmerus dentex

Steindachner and Kner, 1870

Pacific rainbow smelt

Japan Sasaki et al. 2019

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Osmeriformes Osmeridae Osmerus eperlanus (Linnaeus, 1758)

rainbow smelt Baltic Sea, Finland, Tvärminne

Forssell 1904

Osmeriformes Osmeridae Osmerus eperlanus (Linnaeus, 1758)

rainbow smelt Baltic Sea, Finland, Tvärminne

Forssell 1905

Osmeriformes Osmeridae Osmerus eperlanus (Linnaeus, 1758)

rainbow smelt Germany Lühe 1911

Osmeriformes Osmeridae Osmerus eperlanus (Linnaeus, 1758)

rainbow smelt Sweden Lundström 1942 Osmeriformes Osmeridae Osmerus eperlanus

(Linnaeus, 1758)

rainbow smelt Europe Van Cleave 1953 Osmeriformes Osmeridae Osmerus eperlanus

(Linnaeus, 1758)

rainbow smelt Baltic Sea, Bothnian Bay

Valtonen and Julkunen 1995 Perciformes Labridae Ctenolabrus rupestris

(Linnaeus, 1758)

Atlantic herring southern Norway, Flødevigen,

Karlsbakk et al. 1996

Perciformes Percidae Gymnocephalus cernua (Linnaeus, 1758) (as Acerina cernua)

ruffe Baltic Sea,

Finland, Tvärminne

Forssell 1905

Perciformes Percidae Gymnocephalus cernua (Linnaeus, 1758) (as Acerina cernua)

ruffe Germany Lühe 1911

Perciformes Percidae Gymnocephalus cernua (Linnaeus, 1758) (as Acerina cernua)

ruffe Sweden Lundström

1942

Perciformes Percidae Gymnocephalus cernua (Linnaeus, 1758) (as Acerina cernua)

ruffe Europe Van Cleave

1953

Perciformes Percidae Gymnocephalus cernua (Linnaeus, 1758) (as Gymnocephalus cernuus)

ruffe Baltic Sea,

Bothnian Bay

Valtonen and Julkunen 1995

Perciformes Percidae Perca fluviatilis Linnaeus, 1758

perch Baltic Sea,

Finland, Tvärminne

Forssell 1905

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Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Perciformes Percidae Perca fluviatilis Linnaeus, 1758

perch Sweden Lundström

1942 Perciformes Percidae Perca fluviatilis

Linnaeus, 1758

perch Europe Van Cleave

1953 Perciformes Percidae Sander lucioperca

(Linnaeus, 1758) (as Stizostedion lucioperca)

zanda, pike-perch Finno-Karelian ASSR

Rumyantsev and Ieshko 1997* Perciformes Zoarcidae Lycodes raridens

Taranetz & Andriashev, 1937

marbled eelpout Arctic Ocean, Bering Sea

Shults and Frost 1988

Perciformes Zoarcidae Zoarces viviparus (Linnaeus, 1758)

eelpout Sweden Lundström

1942 Perciformes Zoarcidae Zoarces viviparus

(Linnaeus, 1758)

eelpout Europe Van Cleave

1953 Perciformes Zoarcidae Zoarces viviparus

(Linnaeus, 1758)

eelpout Baltic Sea, Estonia, Riga Bay

Vismanis et al. 1980*

Perciformes Zoarcidae Zoarces viviparus (Linnaeus, 1758)

eelpout Baltic Sea, Germany, Lübecker Bay

Zander 1991

Pleuronectiformes Pleuronectidae Hippoglossoides platessoides (Fabricius, 1780) (as

Hippoglossoides platessoides limandoides)

American plaice Arctic Ocean, Greenland/ Icelandic Zone, Icelandic coast

Olafsdottir 1999

Pleuronectiformes Pleuronectidae Limanda limanda (Linnaeus, 1758) (as Pleuronectes limanda)

common dab Sweden Lundström

1942

Pleuronectiformes Pleuronectidae Limanda limanda (Linnaeus, 1758) (as Pleuronectes limanda)

common dab Europe Van Cleave

1953

Pleuronectiformes Pleuronectidae Platichthys flesus (Linnaeus, 1758)

flounder Baltic Sea, Finland, Tvärminne

Forssell 1905

Pleuronectiformes Pleuronectidae Platichthys flesus (Linnaeus, 1758)

(15)

Pleuronectiformes Pleuronectidae Platichthys flesus (Linnaeus, 1758)

flounder Sweden Lundström

1942 Pleuronectiformes Pleuronectidae Platichthys flesus

(Linnaeus, 1758)

flounder Sweden Van Cleave

1953 Pleuronectiformes Pleuronectidae Pleuronectes platessa

Linnaeus, 1758

European plaice Europe Van Cleave 1953 Pleuronectiformes Pleuronectidae Pseudopleuronectes

americanus Walbaum, 1792

winter flounder Atlantic, Canada

Arai 1989

Pleuronectiformes Scophthalmidae Scophthalmus maximus (Linnaeus, 1758) (as Rhombus maximus)

turbot Baltic Sea,

Finland, Tvärminne

Forssell 1904

Pleuronectiformes Scophthalmidae Scophthalmus maximus (Linnaeus, 1758) (as Rhombus maximus)

turbot Baltic Sea,

Finland, Tvärminne

Forssell 1905

Pleuronectiformes Scophthalmidae Scophthalmus maximus (Linnaeus, 1758) (as Rhombus maximus)

turbot Germany Lühe 1911

Pleuronectiformes Scophthalmidae Scophthalmus maximus (Linnaeus, 1758) (as Rhombus maximus)

turbot Sweden Lundström

1942

Pleuronectiformes Scophthalmidae Scophthalmus maximus (Linnaeus, 1758) (as Rhombus maximus)

turbot Europe Van Cleave

1953

Salmoniformes Salmonidae Coregonus albula (Linnaeus, 1758)

European cisco Sweden Lundström 1942 Salmoniformes Salmonidae Coregonus albula

(Linnaeus, 1758)

European cisco Europe Van Cleave 1953 Salmoniformes Salmonidae Coregonus fera Jurine,

1825

true fera Europe Van Cleave

1953 Salmoniformes Salmonidae Salmo salar Linnaeus,

1758

salmon European

USSR (CIS)

Rumyantsev et al. 1998* Salmoniformes Salmonidae Salmo trutta Linnaeus,

1758

brown trout, sea trout

European USSR (CIS)

Rumyantsev et al. 1998* Salmoniformes Salmonidae Thymallus thymallus

(Linnaeus, 1758)

grayling Europe Rumyantsev

(16)

Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758)

fourhorn sculpin Baltic Sea, Bothnian Bay

Sinisalo and Valtonen 2003 Scorpaeniformes Cottidae Myoxocephalus

quadricornis (Linnaeus, 1758) (as Cottus quadricornis)

fourhorn sculpin Baltic Sea, Finland, Tvärminne

Forssell 1904

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758) (as Cottus quadricornis)

fourhorn sculpin Baltic Sea, Finland, Tvärminne

Forssell 1905

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758) (as Cottus quadricornis)

fourhorn sculpin Germany Lühe 1911

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758) (as Cottus quadricornis)

fourhorn sculpin Sweden Lundström 1942

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758) (as Cottus quadricornis)

fourhorn sculpin Europe Van Cleave 1953

Scorpaeniformes Cottidae Myoxocephalus scorpius (Linnaeus, 1758)

shorthorn sculpin Baltic Sea, Bothnian Bay

Valtonen and Julkunen 1995 Scorpaeniformes Cottidae Myoxocephalus

scorpius (Linnaeus, 1758)

shorthorn sculpin Baltic Sea, North-eastern Bothnian Bay

Valtonen et al. 2001

Scorpaeniformes Cottidae Myoxocephalus scorpius (Linnaeus, 1758) (as Cottus scorpius)

shorthorn sculpin Sweden Lundström 1942

Scorpaeniformes Cottidae Myoxocephalus scorpius (Linnaeus, 1758) (as Cottus scorpius)

shorthorn sculpin Baltic Sea, Finland, Tvärminne

Forssell 1905

(17)

Scorpaeniformes Cottidae Myoxocephalus scorpius (Linnaeus, 1758) (as Cottus scorpius)

shorthorn sculpin Europe Van Cleave 1953

Scorpaeniformes Cyclopteridae Cyclopterus lumpus Linnaeus, 1758

goldsinny wrasse Sweden Lundström 1942 Scorpaeniformes Cyclopteridae Cyclopterus lumpus

Linnaeus, 1758

goldsinny wrasse Europe Van Cleave 1953 Scorpaeniformes Psychrolutidae Dasycottus setiger

Bean, 1890

spinyhead sculpin Arctic Ocean, Bering Sea

Shults and Frost 1988

Host

order Host family Host species Host

vernacular name

Geographical locality

Reference

Artiodactyla Monodontidae Delphinapterus leucas (Pallas, 1776)

Beluga, Beluga whale, white whale

Bristol Bay, Alaska, Barent Sea, Sea of Okhotsk, North Pacific Ocean, Arctic Krotov and Delyamure 1952*, Delyamure and Kleinenberg 1958*, Neiland 1962, Yablokov et al. 1972* all cited by Measures et al. 1995 Carnivora Odobenidae Odobenus

rosmarus (Linnaeus, 1758)

walrus no area specified Dailey 1975, Raga 1992

Carnivora Otariidae Callorhinus ursinus (Linnaeus, 1758)

Northern fur seal Alaska, Bering Sea, St. Paul Island

Ionita et al. 2008, Kuzmina et al. 2012

Carnivora Otariidae Zalophus californianus (Lesson, 1828)

California sea lion California Van Cleave 1953

Carnivora Phocidae Cystophora cristata (Erxleben, 1777)

hooded seal Europe Van Cleave 1953

Carnivora Phocidae Erignathus barbatus (Erxleben, 1777)

bearded seal Alaska, Arctic Van Cleave 1953

Table 5.

Definitive hosts of

Corynosoma strumosum

. *Source not viewed. ^Found in Host-Parasite

Database NHM London without further reference details.

(18)

Host order

Host family Host species Host

vernacular name

Geographical locality

Reference

Carnivora Phocidae Erignathus barbatus (Erxleben, 1777)

bearded seal Barent Sea, Sea of Okhotsk, Chukchi seas, Eastern Siberian Sea

Popov and Fortunato 1987

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Germany Lühe 1911

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Holland, Germany Aznar et al. 2006

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Atlantic coast, Ireland

O'Neill and Whelan 2002

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Baltic Sea, Finland, Gulf of Bothnia, Åland Island

Nickol et al. 2002

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Baltic Sea, Baltic Proper, Bothnian Sea, Bothnian Bay, Sweden

Leidenberger and Bäcklin 2008

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal North Sea, Germany, Schleswig-Holstein

Waindok et al. 2018

Carnivora Phocidae Halichoerus grypus (Fabricius, 1791)

grey seal Arctic Ocean, Northern Atlantic, European Arctic

Delyamure 1961*, von Sprehn 1966* cited by Raga 1992 Carnivora Phocidae Halichoerus

grypus (Fabricius, 1791)

grey seal Alaska, Arctic Van Cleave 1953

Carnivora Phocidae Histriophoca fasciata (Zimmermann, 1783)

ribbon seal Barent Sea, Sea of Okhotsk, Chukchi seas, Eastern Siberian Sea

Popov and Fortunato 1987

(19)

Carnivora Phocidae Histriophoca fasciata (Zimmermann, 1783) (as Phoca fasciata)

ribbon seal Bering Sea, Alaska

Rausch et al. 1990

Carnivora Phocidae Histriophoca fasciata (Zimmermann, 1783) (as Phoca fasciata)

ribbon seal Bering Sea Shults and Frost 1988

Carnivora Phocidae Pagophilus groenlandicus (Erxleben, 1777)

harp seal Greenland Van Cleave 1953

Carnivora Phocidae Pagophilus groenlandicus (Erxleben, 1777) (as Phoca groenlandica)

harp seal Germany Lühe 1911

Carnivora Phocidae Phoca largha Pallas, 1811

spotted seal Sea of Okhotsk, Japan Sea

Sasaki et al. 2019

Carnivora Phocidae Phoca largha Pallas, 1811

spotted seal Bering Sea, Alaska

Rausch et al. 1990

Carnivora Phocidae Phoca largha Pallas, 1811

spotted seal Bering Sea [On the helminthofauna of largi and harbour seals.]^ Carnivora Phocidae Phoca largha

Pallas, 1811

Larga seal, spotted seal

Barent Sea, Sea of Okhotsk, Chukchi seas, Eastern Siberian Sea

Popov and Fortunato 1987

Carnivora Phocidae Phoca largha Pallas, 1811 Larga seal, spotted seal Russia: Anadyr Gulf Aznar et al. 2006

Carnivora Phocidae Phoca largha Pallas, 1811

spotted seal no area specified [On the helminthofauna of largi and harbour seals.]^

(20)

Host order

Host family Host species Host

vernacular name

Geographical locality

Reference

Carnivora Phocidae Phoca largha Pallas, 1811

spotted seal Japan, Hokkaido, Nemuro Peninsula

Nakaoka et al. 1986*

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Baltic Sea Mühling 1989

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Germany Rudolphi 1802

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Germany Lühe 1911

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Sweden Lundström 1942

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Baltic Sea and North Sea, Germany, Schleswig-Holstein

Waindok et al. 2018

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Alaska, Bering Sea, St. Paul Island

Kuzmina et al. 2012

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Alaska, Arctic Van Cleave 1953

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal USA, California, Monterey Bay

García-Varela et al. 2011

Carnivora Phocidae Phoca vitulina Linnaeus, 1758

harbour seal Pacific Sasaki et al. 2019

Carnivora Phocidae Phoca vitulina Linnaeus, 1758 (as Phoca vitulina stejnegeri J.A.Allen, 1902)

Kuril harbour seal Japan, Hokkaido, Erimo Cape

Kaimoto et al. 2018

Carnivora Phocidae Phoca vitulina Linnaeus, 1758 (as Phoca vitulina stejnegeri J.A.Allen, 1902)

Kuril harbour seal Japan, Hokkaido, Nemuro Peninsula

(21)

Carnivora Phocidae Phoca vitulina richardii (Gray, 1864)

Californian harbour seal

California Van Cleave 1953

Carnivora Phocidae Phoca vitulina richardii (Gray, 1864)

Pacific harbor seal Washington, Northern Pacific

Dailey and Fallace 1989

Carnivora Phocidae Pusa caspica (Gmelin, 1788)

Caspian seal Iran, Mazandaran Province, Ramsar City

Amin et al. 2011

Carnivora Phocidae Pusa caspica, (Gmelin, 1788) (as Phoca caspica Gmelin, 1788)

Caspian seal Caspian Sea Kurochkin 1975* cited by Raga 1992

Carnivora Phocidae Pusa hispida saimensis (Nordquist, 1899) Saimaa ringed seal Lake Saimaa, Finland

Valtonen and Helle 1988

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Germany Lühe 1911

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Sweden Lundström 1942

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Baltic Sea, Bothnian Bay, Finland

Helle and Valtonen 1981

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Baltic Sea, Bothnian Bay, Finland

Valtonen and Helle 1988

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Baltic Sea, Bothnian Bay, Finland

Valtonen et al. 2004

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Baltic Sea, Sweden

Leidenberger et al. 2019

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Alaska, Arctic Van Cleave 1953

Carnivora Phocidae Pusa hispida (Schreber, 1775)

ringed seal Barent Sea, Sea of Okhotsk, Chukchi seas, Eastern Siberian Sea

Popov and Fortunato 1987

(22)

Host order

Host family Host species Host

vernacular name

Geographical locality

Reference

Carnivora Phocidae Pusa hispida (Schreber, 1775) (as Phoca foetida)

ringed seal Baltic Sea, Finland, Tvärminne

Forssell 1904

Carnivora Phocidae Pusa hispida (Schreber, 1775) (as Phoca foetida)

ringed seal Baltic Sea, Finland, Tvärminne

Forssell 1905

Cetartiodactyla Phocoeridae Phocoena phocoena (Linnaeus, 1758)

harbour porpoise Pacific Sasaki et al. 2019

Cetartiodactyla Phocoeridae Neophocaena phocaenoides (G. Cuvier, 1829)

finless porpoise Pacific Sasaki et al. 2019

Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Acipenseriformes Acipenseridae Acipenser gueldenstaedti Brandt & Ratzeburg, 1833

Danube sturgeon Caspian Sea, southern part, Sefid-Rud River

Sattari and Mokhayer 2005

Acipenseriformes Acipenseridae Acipenser persicus Borodin, 1897 (as Acipenser gueldenstaedti persicus natio kurensis)

Persian sturgeon Georgia, River Kura

Kurashvili et al. 1980*

Acipenseriformes Acipenseridae Acipenser stellatus Pallas, 1771

starry sturgeon Caspian Sea, southern part, Sefid-Rud River

Sattari and Mokhayer 2005

Acipenseriformes Acipenseridae Acipenser transmontanus Richardson, 1836

white sturgeon British Columbia, Canada

Arai 1989

Acipenseriformes Acipenseridae Huso huso (Linnaeus, 1758)

beluga Caspian Sea,

southern part, Sefid-Rud River Sattari and Mokhayer 2005

Table 6.

Paratenic host records for

Corynosoma strumosum

. *Source not viewed, NA: not available, ^Found

in Host-Parasite Database NHM London without further reference details.

(23)

Anguilliformes Anguillidae Anguilla anguilla (Linnaeus, 1758)

European eel Germany Lühe 1911

Anguilliformes Congridae Conger conger (Linnaeus, 1758)

European conger Sweden Lundström 1942 Anguilliformes Congridae Conger conger

(Linnaeus, 1758)

European conger Europe Van Cleave 1953 Clupeiformes Clupeidae Alosa braschnikowi

(Borodin, 1904) (as Alosa brashnikovi grimmi)

Caspian marine shad

Caspian Sea Ibragimov and Vetchanin 1988*

Clupeiformes Clupeidae Alosa braschnikowi (Borodin, 1904) (as Alosa brashnikovi kisselewitschi)

Caspian marine shad

Caspian Sea Ibragimov and Vetchanin 1988*

Clupeiformes Clupeidae Alosa kessleri (Grimm, 1887) (as Alosa kessleri volgensis)

Caspian anadromous shad

Caspian Sea Izyumova 1977*

Clupeiformes Clupeidae Alosa kessleri (Grimm, 1887) (as Alosa kessleri volgensis)

Caspian anadromous shad

USSR (CIS) Izyumova 1977*

Clupeiformes Clupeidae Clupea harengus Linnaeus, 1758

herring Sweden Lundström

1942 Clupeiformes Clupeidae Clupea harengus

Linnaeus, 1758

herring Europe Van Cleave

1953 Clupeiformes Clupeidae Clupea harengus

Linnaeus, 1758 (as Clupea harengus membras)

herring Baltic Sea, Finland, Tvärminne

Forssell 1905

Clupeiformes Clupeidae Clupea pallasii Valenciennes, 1847

Pacific herring Pacific ocean, Sea of Okhotsk

Sasaki et al. 2019

(24)

Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Clupeiformes Clupeidae Clupea pallasii pallasii

Valenciennes, 1847 (as Clupea harengus pallasii)

Pacific herring Pacific ocean, Canada

Arai 1989

Clupeiformes Clupeidae Clupeonella caspia Svetovidov, 1941 (as Clupeonella delicatula caspia)

Caspian tyulka Caspian Sea Ibragimov 1988*

Clupeiformes Clupeidae Clupeonella cultriventris (Nordmann, 1840) (as Clupea cultriventris)

Black Sea sprat Caspian Sea Shamsi et al. 1998*

Clupeiformes Clupeidae Clupeonella grimmi Kessler, 1877 Caspian sprat, big-eyed kilka Capian Sea, southern part Habibi and Shamsi 2018 Cypriniformes Cyprinidae Abramis brama

(Linnaeus, 1758)

freshwater bream Georgia, River Kura

Kurashvili et al. 1980* Cypriniformes Cyprinidae Alburnus chalcoides

(Güldenstädt, 1772) (as Chalcalburnus chalcoides)

shemaya Georgia, River Kura (USSR)

Kurashvili et al. 1980*

Cypriniformes Cyprinidae Alburnus chalcoides (Güldenstädt, 1772) (as Chalcalburnus chalcoides)

shemaya USSR (CIS) Izyumova

1977*

Cypriniformes Cyprinidae Barbus barbus (Linnaeus, 1758)

barbel USSR (CIS) Izyumova

1977* Cypriniformes Cyprinidae Barbus

brachycephalus caspius Berg, 1914

Caspian barbel River Kura (USSR)

Kurashvili et al. 1980*

Cypriniformes Cyprinidae Carassius carassius (Linnaeus, 1758)

Crucian carp Azerbaijan On the study of helminths of the crucian carp in Azerbaijan fish farms.

(25)

Cypriniformes Cyprinidae Cyprinus carpio Linnaeus, 1758

common carp Georgia, River Kura (USSR)

Kurashvili et al. 1980* Cypriniformes Cyprinidae Leuciscus idus

(Linnaeus, 1758)

orfe, ide Finno-Karelian ASSR

Rumyantsev and Ieshko 1997* Cypriniformes Cyprinidae Pelecus cultratus

(Linnaeus, 1758) sabrefish, razorfish, knife Finno-Karelian ASSR Rumyantsev and Ieshko 1997* Cypriniformes Cyprinidae Rutilus frisii

(Nordmann, 1840) (as Rutilus frisii kutum)

Black Sea roach Georgia, River Kura (USSR)

Kurashvili et al. 1980*

Cypriniformes Cyprinidae Rutilus rutilus (Linnaeus, 1758) (as Rutilus rutilus caspicus natio knipowitschia)

roach Caspian Sea Ibragimov

1989*

Cypriniformes Cyprinidae Vimba vimba (Linnaeus, 1758)

vimba bream Lithuania Rauckis 1988*

Esociformes Esocidae Esox lucius Linnaeus, 1758

pike Baltic Sea,

Finland, Tvärminne

Forssell 1905

Gadiformes Gadidae Boreogadus saida (Lepechin, 1774)

Polar cod Arctic Ocean: USSR zone

Karasev and Mitenev 1993* Gadiformes Gadidae Eleginus gracilis

(Tilesius, 1810)

saffron cod Gertner Bay, Cape of Nyuklya near Magadan Skorobrechova and Nikishin 2014

Gadiformes Gadidae Eleginus gracilis (Tilesius, 1810)

saffron cod Japan, Shikotan Island

Zhukov 1960*

Gadiformes Gadidae Eleginus nawaga (Walbaum, 1792) (as Eleginus navaga)

Arctic cod Arctic Ocean: USSR zone

Karasev and Mitenev 1993*

(26)

Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Gadiformes Gadidae Gadus

chalcogrammus Pallas, 1814 (as Theragra chalcogramma)

Alaska pollock Bering Sea Shults and Frost 1988

Gadiformes Gadidae Gadus

chalcogrammus Pallas, 1814 (as Theragra chalcogramma)

Alaska pollock Bering Sea Avdeev and Avdeev 1998*

Gadiformes Gadidae Gadus

chalcogrammus Pallas, 1814 (as Theragra chalcogramma)

Alaska pollock Pacific ocean, Canada

Arai 1989

Gadiformes Gadidae Gadus

macrocephalus Tilesius, 1810

Pacific cod Pacific coast, USA and Canada

Van Cleave 1953

Gadiformes Gadidae Gadus morhua

Linnaeus, 1758 (as Gadus callarias)

cod Sweden Lundström

1942

Gadiformes Gadidae Gadus morhua

Linnaeus, 1758 (as Gadus callarias)

cod Europe Van Cleave

1953

Gadiformes Gadidae Melanogrammus

aeglefinus (Linnaeus, 1758)

haddock Atlantic, Canada

Arai 1989

Gadiformes Lotidae Lota lota (Linnaeus, 1758)

burbot Baltic Sea,

Bothnian Bay

Valtonen and Julkunen 1995 Gadiformes Lotidae Lota lota (Linnaeus,

1758) (as Lota vulgaris)

burbot Baltic Sea,

Finland, Tvärminne

Forssell 1905

Gadiformes Lotidae Lota lota (Linnaeus, 1758) (as Lota vulgaris)

burbot Germany Lühe 1911

Gadiformes Lotidae Lota lota (Linnaeus, 1758) (as Lota vulgaris)

burbot Sweden Lundström

(27)

Gadiformes Lotidae Lota lota (Linnaeus, 1758) (as Lota vulgaris)

burbot Europe Van Cleave

1953

Gadiformes Merlucciidae Macruronus novaezelandiae (Hector, 1871) whiptail, tailed hake, Patagonian whiphake, New Zealand whiptail, blue hake, blue grenadier

New Zealand coast

Klimpel et al. 2001*

Gadiformes Merlucciidae Merluccius capensis Castelnau, 1861

whiting, stockfish, South African whiting, shallow-water Cape hake, shallow water hake, hake, Cape hake

no area specified

Parukhin 1989*

Gasterosteiformes Gasterosteidae Gasterosteus aculeatus Linnaeus, 1758 three-spined stickleback Sweden Lundström 1942

Gasterosteiformes Gasterosteidae Gasterosteus aculeatus Linnaeus, 1758

three-spined stickleback

Europe Van Cleave 1953

Gasterosteiformes Gasterosteidae Gasterosteus aculeatus Linnaeus, 1758 three-spined stickleback Caspian Sea, southeast part, Gomishan Lagoon Niksirat et al. 2006

Lophiiformes Lophiidae Lophius piscatorius Linnaeus, 1758

angler Baltic Sea Mühling 1989

Lophiiformes Lophiidae Lophius piscatorius Linnaeus, 1758

angler Sweden Lundström

1942 Lophiiformes Lophiidae Lophius piscatorius

Linnaeus, 1758

angler Europe Van Cleave

1953 Lophiiformes Lophiidae Lophius piscatorius

Linnaeus, 1758 monkfish, angler-fish no area specified Parukhin 1989*

Ophidiiformes Ophidiidae Hoplobrotula gnathopus (Regan, 1921)

false kinglip no area specified

(28)

Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Osmeriformes Osmeridae Hypomesus japonicus (Brevoort, 1856)

Japanese smelt Pacific ocean Sasaki et al. 2019

Osmeriformes Osmeridae Hypomesus japonicus (Brevoort, 1856)

Japanese smelt Japan, Shikotan Island

Zhukov 1960*

Osmeriformes Osmeridae Hypomesus japonicus (Brevoort, 1856)

Japanese smelt Japan, Hokkaido, Erimo Cape

Araki and Machida 1987*

Osmeriformes Osmeridae Hypomesus japonicus (Brevoort, 1856) (as

Hypomesus preiosus japonicus)

Japanese smelt Japan, Hokkaido

Nagasawa et al. 1989*

Osmeriformes Osmeridae Hypomesus olidus (Pallas, 1814)

pond smelt Nagaevo and Gertner Bay, Magadan, Russia, Sea of Okhotsk Skorobrechova and Nikishin 2011

Osmeriformes Osmeridae Hypomesus olidus (Pallas, 1814)

pond smelt Japan, Shikotan Island

Zhukov 1960*

Osmeriformes Osmeridae Osmerus dentex Steindachner and Kner, 1870 Pacific rainbow smelt Japan, Sea of Okhotsk, Pacific ocean Sasaki et al. 2019

Osmeriformes Osmeridae Osmerus eperlanus (Linnaeus, 1758)

rainbow smelt Sweden Lundström 1942 Osmeriformes Osmeridae Osmerus eperlanus

(Linnaeus, 1758)

rainbow smelt Europe Van Cleave 1953 Osmeriformes Osmeridae Osmerus eperlanus

(Linnaeus, 1758)

rainbow smelt Baltic Sea, Bothnian Bay

Valtonen and Julkunen 1995 Osmeriformes Osmeridae Osmerus mordax

dentex Steindachner & Kner, 1870

rainbow smelt Nagaevo and Gertner Bay, Magadan, Russia, Sea of Okhotsk Skorobrechova and Nikishin 2011

(29)

Osmeriformes Osmeridae Spirinchus lanceolatus (Hikita, 1913)

Sishamo smelt Japan Hokkaido, Mukawa River

Fujita 1921*

Osmeriformes Osmeridae Spirinchus lanceolatus (Hikita, 1913) (as Osmerus lanceolatus)

Shishamo, willow leaf fish

Japan Van Cleave

1953

Perciformes Gobiidae Benthophilus stellatus (Sauvage, 1874)

Stellate tadpole-goby

Caspian Sea Ibragimov 1988*

Perciformes Gobiidae Gobius Linnaeus, 1758

goby Caspian Sea Ibragimov

1988* Perciformes Gobiidae Neogobius fluviatilis

(Pallas, 1814) (as Gobius fluviatilis)

monkey goby Caspian Sea Ibragimov 1988*

Perciformes Gobiidae Neogobius fluviatilis (Pallas, 1814) (as Neogobius fluviatilis pallasi (Pallas, 1814)

monkey goby Caspian Sea, south-eastern part

Pazooki et al. 2011

Perciformes Gobiidae Neogobius melanostomus (Pallas, 1814)

round goby Caspian Sea Ibragimov 1988*

Perciformes Gobiidae Ponticola bathybius (Kessler, 1877) (as Neogobius bathybius ) NA Capian Sea, south-eastern part Pazooki et al. 2011

Perciformes Gobiidae Ponticola gorlap (Iljin, 1949) (as Neogobius kessleri gorlap) Caspian bighead goby Capian Sea, south-eastern part Pazooki et al. 2011

Perciformes Gobiidae Ponticola kessleri (Günther, 1861) (as Neogobius kessleri)

bighead goby Caspian Sea Ibragimov 1988*

Perciformes Hexagrammidae Hexagrammos octogrammus (Pallas, 1814)

masked greenling Sea of Okhotsk

Skorobrechova 2009* cited in Skorobrechova et al. 2011

(30)

Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Perciformes Mugilidae Chelon auratus (Risso, 1810) (as Mugil auratus)

golden grey mullet

Caspian Sea Ibragimov 1988*

Perciformes Mugilidae Chelon auratus (Risso, 1810) (as Mugil auratus)

leaping mullet Caspian Sea Ibragimov 1988*

Perciformes Percidae Gymnocephalus cernua (Linnaeus, 1758) (as Gymnocephalus cernuus)

ruffe Baltic Sea,

Bothnian Bay

Valtonen and Julkunen 1995

Perciformes Percidae Perca fluviatilis Linnaeus, 1758

perch Baltic Sea,

Finland, Tvärminne

Forssell 1905

Perciformes Percidae Perca fluviatilis Linnaeus, 1758

perch Germany Lühe 1911

Perciformes Percidae Perca fluviatilis Linnaeus, 1758

perch Europe Van Cleave

1953 Perciformes Percidae Sander lucioperca

(Linnaeus, 1758) (as Lucioperca lucioperca) zander, pike-perch Georgia, River Kura (USSR) Kurashvili et al. 1980*

Perciformes Percidae Sander lucioperca (Linnaeus, 1758) (as Lucioperca lucioperca)

zander, pike-perch

USSR (CIS) Izyumova 1977*

Perciformes Percidae Sander lucioperca (Linnaeus, 1758) (as Stizostedion lucioperca) zander, pike-perch Finno-Karelian ASSR Rumyantsev and Ieshko 1997*

Perciformes Sciaenidae Umbrina roncador Jordan & Gilbert, 1882

yellowfin croaker Catalina Island, California

Van Cleave 1953

Perciformes Sparidae Pagrus pagrus (Linnaeus, 1758) red porgy, Couch's sea-bream, common seabream no area specified Parukhin 1966*

(31)

Perciformes Sparidae Sparus heterodus Peters, 1877

NA no area

specified

Parukhin 1966*

Perciformes Sphyraenidae Sphyraena barracuda (Edwards, 1771)

great barracuda no area specified

Parukhin 1966*

Perciformes Stichaeidae Pholidapus dybowskii

(Steindachner, 1880)

stichaeid fish Japan, Shikotan Island

Zhukov 1960*

Perciformes Trachinidae Trachinus draco Linnaeus, 1758

weever Baltic Sea Mühling 1989

Perciformes Trachinidae Trachinus draco Linnaeus, 1758

weever Sweden Lundström

1942 Perciformes Trachinidae Trachinus draco

Linnaeus, 1758

weever Germany Lühe 1911

Perciformes Trachinidae Trachinus draco Linnaeus, 1758

weever Europe Van Cleave

1953 Perciformes Trichodontidae Arctoscopus

japonicus (Steindachner, 1881) Japanese sandfish Japan, Shikotan Island Zhukov 1960*

Perciformes Zoarcidae Lycodes raridens Taranetz & Andriashev, 1937

marbled eelpout Bering Sea Shults and Frost 1988

Perciformes Zoarcidae Hadropareia middendorffii Schmidt, 1904 NA Sea of Okhosk Skorobrechova 2010* cited in Skorobrechova et al. 2011 Perciformes Zoarcidae Zoarces viviparus

(Linnaeus, 1758)

eelpout Baltic Sea, Polen, Gdansk Bight

Markowski 1938

Perciformes Zoarcidae Zoarces viviparus (Linnaeus, 1758)

eelpout Sweden Lundström

1942 Perciformes Zoarcidae Zoarces viviparus

(Linnaeus, 1758)

eelpout Europe Van Cleave

1953 Perciformes Zoarcidae Zoarces viviparus

(Linnaeus, 1758)

eelpout Baltic Sea, Germany, Salzhaff

Reimer and Walter 1998

(32)

Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Pleuronectiformes Pleuronectidae Atheresthes stomias (Jordan & Gilbert, 1880) arrow-tooth flounder British Columbia Wierzbicka and Piasecki 1998*

Pleuronectiformes Pleuronectidae Eopsetta jordani (Lockingston, 1879)

petrale sole Canada Arai 1989

Pleuronectiformes Pleuronectidae Hippoglossoides platessoides (Fabricius, 1780) (as Hippoglossoides platessoides limandoides) sand-dab, long rough dab, Canadian plaice, American plaice Arctic Ocean, Greenland/ Icelandic zone Olafsdottir 1999

Pleuronectiformes Pleuronectidae Hippoglossus stenolepis (Schmidt, 1904)

Pacific halibut Northern Pacific

Blaylock et al. 1998*

Pleuronectiformes Pleuronectidae Lepidopsetta bilineata (Ayres, 1855)

rock sole Pacific coast, USA and Canada

Van Cleave 1953

Pleuronectiformes Pleuronectidae Lepidopsetta bilineata (Ayres, 1855)

rock sole Pacific ocean, Canada

Arai 1989

Pleuronectiformes Pleuronectidae Limanda aspera (Pallas, 1814)

yellowfin sole Nagaevo and Gertner Bay, Magadan, Russia, Sea of Okhotsk Skorobrechova and Nikishin 2011

Pleuronectiformes Pleuronectidae Limanda limanda (Linnaeus, 1758) (as Pleuronectes limanda)

common dab Europe Van Cleave

1953

Pleuronectiformes Pleuronectidae Platichthys flesus (Linnaeus, 1758)

flounder Baltic Sea, Finland, Tvärminne

Forssell 1905

Pleuronectiformes Pleuronectidae Platichthys flesus (Linnaeus, 1758)

flounder Germany Lühe 1911

Pleuronectiformes Pleuronectidae Platichthys flesus (Linnaeus, 1758)

flounder Sweden Lundström

(33)

Pleuronectiformes Pleuronectidae Platichthys flesus (Linnaeus, 1758)

perch Sweden Lundström

1942 Pleuronectiformes Pleuronectidae Platichthys flesus

(Linnaeus, 1758) (as Platessa flesus)

flounder Baltic Sea Mühling 1989

Pleuronectiformes Pleuronectidae Platichthys flesus (Linnaeus, 1758) (as Pleuronectes flesus)

flounder Baltic Sea, Finland, Tvärminne

Forssell 1905

Pleuronectiformes Pleuronectidae Platichthys flesus (Linnaeus, 1758) (as Pleuronectes flesus)

flounder Europe Van Cleave

1953

Pleuronectiformes Pleuronectidae Platichthys stellatus (Pallas, 1787)

starry flounder Japan, Shikotan Island

Zhukov 1960*

Pleuronectiformes Pleuronectidae Platichthys stellatus (Pallas, 1787)

starry flounder Pacific coast, USA and Canada

Van Cleave 1953

Pleuronectiformes Pleuronectidae Platichthys stellatus (Pallas, 1787)

starry flounder Pacific ocean, Canada

Arai 1989

Pleuronectiformes Pleuronectidae Pleuronectes obscurus Herzenstein, 1890

black plaice Japan, Shikotan Island

Zhukov 1960*

Pleuronectiformes Pleuronectidae Reinhardtius hippoglossoides (Walbaum, 1792)

turbot Atlantic,

Canada

Arai 1989

Pleuronectiformes Pleuronectidae Verasper moseri Jordan & Gilbert, 1898

barfin flounder Japan, Hokkaido, Erimo Cape

Araki and Machida 1987*

Pleuronectiformes Scophthalmidae Scophthalmus maximus (Linnaeus, 1758) (as Rhombus maximus)

trubot Baltic Sea,

Finland, Tvärminne

Forssell 1904

Pleuronectiformes Scophthalmidae Scophthalmus maximus (Linnaeus, 1758) (as Rhombus maximus)

trubot Baltic Sea,

Finland, Tvärminne

(34)

Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Pleuronectiformes Scophthalmidae Scophthalmus maximus (Linnaeus, 1758) (as Rhombus maximus)

trubot Germany Lühe 1911

Pleuronectiformes Scophthalmidae Scophthalmus maximus (Linnaeus, 1758) (as Rhombus maximus)

trubot Sweden Lundström

1942

Pleuronectiformes Scophthalmidae Scophthalmus maximus (Linnaeus, 1758) (as Rhombus maximus)

trubot Europe Van Cleave

1953

Salmoniformes Hemitripteridae Hemitripterus villosus (Pallas, 1814)

shaggy sea raven Japan, Hokkaido, Nemuro Peninsula

Nakaoka et al. 1986*

Salmoniformes Salmonidae Coregonus fera Jurine, 1825

true fera Europe Van Cleave

1953 Salmoniformes Salmonidae Coregonus lavaretus

(Linnaeus, 1758)

lavaret Europe Van Cleave

1953 Salmoniformes Salmonidae Oncorhynchus

gorbuscha (Walbaum, 1792)

pink salmon Pacific ocean, Canada

Arai 1989

Salmoniformes Salmonidae Oncorhynchus gorbuscha (Walbaum, 1792)

pink salmon Sakhalin Island

Vyalova 2003*

Salmoniformes Salmonidae Oncorhynchus keta (Walbaum, 1792)

chum salmon Sakhalin Island

Vyalova 2003*

Salmoniformes Salmonidae Oncorhynchus masou (Brevoort, 1856) (as Oncorhynchus masu )

masu salmon Russia Ermolenko et al. 1989*

Salmoniformes Salmonidae Oncorhynchus nerka (Walbaum, 1792)

sockeye salmon Pacific ocean, Canada

Arai 1989

Salmoniformes Salmonidae Salmo salar Linnaeus, 1758

salmon European

USSR (CIS)

Rumyantsev et al. 1998*

(35)

Salmoniformes Salmonidae Salmo trutta Linnaeus, 1758 trout European USSR (CIS) Rumyantsev et al. 1998* Salmoniformes Salmonidae Salvelinus alpinus

alpinus (Linnaeus, 1758)

Arctic charr Canada, Quebec, Ungava Bay

Desdevises et al. 1998*

Salmoniformes Salmonidae Salvelinus alpinus alpinus (Linnaeus, 1758) (as Salvelinus alpinus)

Arctic charr River Chaun [On the helminthofauna of fish of the River Chaun.]^ Salmoniformes Salmonidae Salvelinus

leucomaenis leucomaenis (Pallas, 1814) white-spotted charr Japan, Shikotan Island Zhukov 1960*

Salmoniformes Salmonidae Salvelinus leucomaenis leucomaenis (Pallas, 1814) (as Salvelinus leucomaenis) white-spotted charr Japan, Hokkaido Nagasawa et al. 1989*

Salmoniformes Salmonidae Salvelinus malma (Walbaum, 1792)

dolly varden Russia Ermolenko 1994* Salmoniformes Salmonidae Salvelinus malma

(Walbaum, 1792)

dolly varden Russia Ermolenko et al. 1989* Salmoniformes Salmonidae Thymallus thymallus

(Linnaeus, 1758) grayling European waters of NE Atlantic and Mediterranean Rumyantsev et al. 1999*

Scorpaeniformes Cottidae Gymnocanthus galeatus Bean, 1881

armoured sculpin Bering Sea Shults and Frost 1988 Scorpaeniformes Cottidae Gymnocanthus

herzensteini Jordan & Starks, 1904

staghorn sculpin Japan, Hokkaido, Erimo Cape

Araki and Machida 1987*

Scorpaeniformes Cottidae Leptocottus armatus Girard, 1854 Pacific staghorn sculpin Pacific coast, USA and Canada Van Cleave 1953

Scorpaeniformes Cottidae Myoxocephalus brandtii

(Steindachner, 1867)

snowy sculpin Japan, Shikotan Island

(36)

Host order Host family Host species Host

vernacular name

Geographical locality

Reference

Scorpaeniformes Cottidae Myoxocephalus polyacanthocephalus (Pallas, 1814) (as Ainocottus ensiger)

great sculpin Japan, Hokkaido, Erimo Cape

Araki and Machida 1987*

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758)

fourhorn sculpin Baltic Sea, Bothnian Bay

Sinisalo and Valtonen 2003

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758) (as Cottus quadricornis)

fourhorn sculpin Sweden Lundström 1942

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758) (as Cottus quadricornis)

fourhorn sculpin Baltic Sea, Finland, Tvärminne

Forssell 1904

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758) (as Cottus quadricornis)

fourhorn sculpin Baltic Sea, Finland, Tvärminne

Forssell 1905

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758) (as Cottus quadricornis)

fourhorn sculpin Germany Lühe 1911

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758) (as Cottus quadricornis)

fourhorn sculpin Sweden Lundström 1942

Scorpaeniformes Cottidae Myoxocephalus quadricornis (Linnaeus, 1758) (as Cottus quadricornis)

fourhorn sculpin Europe Van Cleave 1953

Scorpaeniformes Cottidae Myoxocephalus scorpius (Linnaeus, 1758)

fourhorn sculpin Baltic Sea, Bothnian Bay

Valtonen and Julkunen 1995

Scorpaeniformes Cottidae Myoxocephalus scorpius (Linnaeus, 1758)

fourhorn sculpin Baltic Sea, North-eastern Bothnian Bay

Valtonen et al. 2001

Figure

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